BLUMEA 49: 69–81

Published on 3 May 2004

doi: 10.3767/000651904X486197

REVIEW OF THE GENUS SOLENA (CUCURBITACEAE) W.J.J.O. DE WILDE & B.E.E. DUYFJES Nationaal Herbarium Nederland, University Leiden branch P.O. Box 9514, 2300 RA Leiden, The Netherlands; e-mail: [email protected] SUMMARY

The genus Solena Lour. is redefined, and its 3 species and 1 subspecies are keyed out and described: S. amplexicaulis occurs in S India, S. heterophylla, with 2 subspecies is widespread, but does not occur in S India, and S. umbellata (Klein ex Willd.) W.J. de Wilde & Duyfjes, comb. nov., occurs in S India and Sri Lanka. The status of S. heterophylla subsp. napaulensis (Ser.) W.J. de Wilde & Duyfjes, comb. & stat. nov. needs further study as this subspecies is very similar to the more widespread subsp. heterophylla, but it is distinct by sigmoid thecae. Solena amplexicaulis is taxonomically more distinct from the other two species of Solena. In southern India Kedrostis foetidissima much resembles Solena in general habit, especially by the more or less amplexicaul leaves, and therefore it is discussed and keyed out against Solena. Key words: Cucurbitaceae, Solena, SE Asia.

INTRODUCTION

In the past the genus Solena Lour. was regarded to consist of only one widespread variable species, S. amplexicaulis (Lam.) Gandhi, ranging from NE Afghanistan through India and Sri Lanka, eastward to eastern and southern China, and into West Malesia (Jeffrey, 1980b). However, for China (Wu, 1984; Lu & Zhang, 1986; Chen, 1995) a second species, S. delavayi (Cogn.) C.Y. Wu, was accepted. Examining Solena over its whole area we found that 3 species can be recognised: 1) S. amplexicaulis in S India; 2) S. umbellata in S India and Sri Lanka; and 3) S. heterophylla with two subspecies: subsp. heterophylla, variable in general appearance and widely distributed from NE Afghanistan eastward, and subsp. napaulensis (synonym S. delavayi) in the area of N India, east to China. The latter occurs in mountainous areas; it is likewise variable in general habit and very similar to the type subspecies, but distinct by its obvious sigmoid thecae. According to the above cited Chinese authors, S. delavayi is further distinct by being monoecious and by having deeply 5-lobed (not 3-lobed or subentire) leaves, but we doubt whether these additional characters hold true unambiguously. The remaining three Solena taxa have straight or curved, never S-shaped, thecae. As the two character states for the thecae, straight or curved versus sigmoid, is a strong argument in the distinction of genera in Cucurbitaceae, it is surprising that, because of the absence of additional characters, one has to admit both character states within one single species S. heterophylla. For the sake of convenience we accept the plants with sigmoid thecae as a separate subspecies, subsp. napaulensis. This means that within the genus Solena, apparently as a unique exception, species occur with straight and © 2004 Nationaal Herbarium Nederland, Leiden University branch

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curved thecae, as well as with sigmoid thecae. A comparable exception of a strong genus character, variable within one genus, viz. stamens free versus stamens connate, is known in Neoalsomitra (De Wilde & Duyfjes, 2003). Members of the genus Solena are easily recognised in the field by their comparatively short petioles, with the blade more or less embracing the stem. In southern India and Sri Lanka this character has led to incidental confusion of sterile S. amplexicaulis and S. umbellata with Kedrostis foetidissima (synonym K. rostrata); all three species superficially look very much alike, because of short-petioled leaves. Furthermore, Kedrostis foetidissima and S. amplexicaulis both have rather similar beaked fruits. Therefore, K. foetidissima is included in the key to the species. The genus Kedrostis contains several species in Madagascar and Africa and two species in Asia. The Asian species of Kedrostis are distinct from Solena e.g. in the male flowers by short filaments, inserted in the throat of the receptacle-tube, by the connective produced beyond the anther-thecae, and the apparent absence of a disc (as it is adnate to the receptacle-tube). Asian Kedrostis can also be confounded with Corallocarpus, which differs in having anthers without produced connective, and with fruit opening in an operculate manner, with a ring-shaped seam at base. The fruits of most Kedrostis and of Solena are indehiscent. The petioles of Corallocarpus and Kedrostis (except K. foetidissima) are comparatively long. Solena is characterised by basally inserted stamens, and a conspicuous glabrous or hairy disc, which is sometimes cup-shaped and deeply lobed, but mostly consists of thickish strap-shaped lobes, at the base of the receptacle-tube in both sexes. These lobes are not to be regarded as pistillode or staminodes, because in female flowers they are present around the base of the style, beside the slightly higher-up inserted small staminodes. In male flowers they are inserted between the stamens and there is no central remnant of a style. SOLENA Solena Lour. (1790) 514; Keraudren (1975) 62. — Melothria sect. Solena (Lour.) Cogn. (1881) 604, p.p.; Pax (1889) 15, p.p.; Cogn. (1916) 104, p.p., for M. heterophylla, M. delavayi and M. amplexicaulis only. — Type: Solena heterophylla Lour. Karivia Arn. (1840) 50; (1841) 275; Miq. (1855) 660. — Type: Karivia amplexicaulis (Lam.) Arn.

Herbaceous or at base subwoody climbers, 2–6 m tall, with perennial rootstock or tubers; monoecious or dioecious. Leaves simple, extremely variable in shape; petiole short, rendering the leaves frequently semi-amplexicaul, and with the basal lobes often overlapping. Tendrils simple. Probract minute, linear, often caducous or absent. Flowers white or cream coloured. Male flowers in sessile or peduncled condensed racemes. Pedicels suberect, persistent, often with bracts at base or inserted about halfway. Receptacle-tube cup-shaped. Sepals 5, minute. Petals 5, free, valvate-induplicate, small, hairy. Stamens 3, two 2-thecous, one 1-thecous. Filaments free, long, inserted at base of the receptacle-tube. Anthers without or with swollen connective, not produced beyond the anther-thecae; thecae either erect and straight, or oblique and (slightly) curved, or sigmoid. Disc 3- (or 4-)lobed, conspicuous, carnose. Female flowers single, solitary or at base of male raceme; perianth resembling male. Ovary narrowly ellipsoid, glabrous or hairy; ovules few or rather many. Staminodes 3 (or 4), inserted on the receptacle. Disc



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a deeply lobed cup or consisting of 3(–5) lobes, at base of style. Fruit fleshy, glabrous, hairy or scabrous. Seeds few or rather many, little compressed or nearly globose. An Asian genus of 3 species and 1 subspecies of variable appearance, mainly due to variable leaf shape. KEY TO SOLENA AND KEDROSTIS FOETIDISSIMA

a. Plant hairy; monoecious. Male flowers without disc; filaments shorter than anthers, inserted in the throat of the receptacle-tube. [Thecae straight; connective produced beyond the anther-thecae.] Fruit beaked, with few pear-shaped seeds. — Africa, S India, Sri Lanka . . . . . . . . . . . . . . . . . . . . . Kedrostis foetidissima (Jacq.) Cogn. b. Plant hairy, (sub)glabrous, or scabrous; monoecious or dioecious. Male flowers with disc consisting of lobes; filaments longer than anthers, inserted at base of the receptacle-tube. Fruit beaked or unbeaked, with few or several globose or somewhat flattened seeds. — Asia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Solena KEY TO THE SPECIES OF SOLENA

1a. Dioecious. Male bracts c. 1 mm long or less, or absent, at base of pedicel, persistent or caducous. Anther-thecae straight, vertical; connective narrow. Ovary apically narrowed into a long neck. Fruit at apex narrowed into a long beak; seeds 1–4 per fruit, c. 9 mm long, warty. — S India . . . . . . . . . . . . . . . . . . . 1. S. amplexicaulis b. Monoecious or dioecious. Male bracts 2–8 mm long, inserted about median on the pedicel, persistent or caducous. Anther-thecae curved or sigmoid, subvertical or transverse; connective swollen. Ovary apically not narrowed. Fruit at apex acute; seeds 5–40 per fruit, 6–7 mm long, smooth or finely warted. — S, N & NE India, Sri Lanka, east to S China, Indochina, Malesia (Peninsular Malaysia, Java) . . . . 2 2a. Male bracts terete, carnose, 2–3 mm long, without glands. Petals with hairs and with gland hairs. Ovary glabrous. Fruit glabrous, ± 9-lined or angular. Seeds with broad, finely warty margin. — S India, Sri Lanka . . . . . . . . . . . . 3. S. umbellata b. Male bracts flat or terete, membranous or slightly carnose, 3–8 mm long, without or with glands. Petals hairy, but without gland hairs. Ovary hairy. Fruit scabrous hairy, not lined nor angular. Seeds with faint smooth margin. — Widespread from NE Afghanistan, N India, to S China, southeast to Peninsular Malaysia, Java . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2. S. heterophylla 1. Solena amplexicaulis (Lam.) Gandhi — Fig. 1d–i Solena amplexicaulis (Lam.) Gandhi (1976) 179; C. Jeffrey (1980a) 801, p.p.; K.M. Matthew (1982) plate 303; (1983) 651, p.p. — Bryonia amplexicaulis Lam. (1785) 496; Ser. (1828) 306; Wight & Arn. (1834) 346; Wight (1841) t. 502. — Karivia amplexicaulis (Lam.) Arn. (1841) 275; M. Roem. (1846) 46. — Melothria amplexicaulis (Lam.) Cogn. (1881) 621; (1916) 125; Gamble (1919) 539; Chakrav. (1959) 146. — Type: Sonnerat s.n. (P), S India.

Extensive vine to 6 m, almost glabrous; dioecious. Leaves ovate or (narrowly) elliptic, 2.5–8 by 2–5 cm, ± 5-angular or deeply and narrowly hastately lobed, base deeply

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a

e

g

f

d

b

h

c

i

Fig. 1. a–c: Solena heterophylla Lour. subsp. napaulensis (Ser.) W.J. de Wilde & Duyfjes. a. Male flower; b, c. female flower. — d – i: Solena amplexicaulis (Lam.) Gandhi. d. Habit of male plant; e. male flower; f, g. female flower; h. fruit; i. seed (a: Ohasi et al. 773353; b, c: Thomson s. n., HLB 901.288.174; d: Matthew, Britto & Rani RHT 28388; e: Matthew & Arockissamy RHT 2562; f, g: Matthew RHT 72949; h, i: Matthew & Rajendran RHT 44545).



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cordate (or hastate), apex rounded or acute, often c. 2 mm mucronate, margin remotely dentate, basal nerves (3–)5, reticulation coarse, not very distinct, glands abaxially few, scattered; petiole 0.2–0.4 cm long. Probract terete, linear or club-shaped, ± carnose, 2–3 mm long, apex subtruncate, shortly-roughly hairy. Male inflorescences: peduncle 10–20 mm long; raceme condensed, 5–10-flowered; pedicels 2–5(–7) mm long, without or with only an indication of a fugacious linear bract at base, c. 1 mm long, and in addition collateral with the peduncle a solitary c. 10 mm long pedicelled flower. Male flowers: glabrous except petals; pedicel articulate 1–2 mm below apex; bracts absent or linear, 1–2 mm long, fugacious; receptacle-tube 3–4 by 2.5–3 mm; sepals c. 0.5 mm long; petals long-triangular, 1.5–2 by 1–1.5 mm, hairy at both sides; filaments erect, 2–2.5 mm long, inserted at the base of the receptacle-tube; anthers mutually closely appressed (not connate), ellipsoid, shallowly notched at apex, 1.8–2 by c. 1 mm; thecae straight, vertical; connective not produced beyond the anther-thecae, narrow, glabrous; disc lobes 3, glabrous, c. 2 mm long. Female flowers: solitary, glabrous (except petals); pedicel 2–2.5 mm long; ovary consisting of an ovoid, c. 3 by 2 mm, fertile part, at apex narrowed into a 6–8 mm long, less than 1 mm thick ‘neck’; receptacle-tube, c. 4 by 4 mm; sepals as in male; petals larger than in male, ovate-elliptic, 3.5–4 by c. 2.5 mm, with rounded apex, 3(–5) parallel-veined, minutely pale hairy; style c. 4 mm long, glabrous; stigma consisting of 3 conspicuous frondose (deeply dissected) lobes, together c. 3.5 mm diam.; disc lobes (3–)5, glabrous, c. 1 mm long; staminodes narrowly elliptic, c. 0.5 mm long, inserted slightly above the base in the receptacle-tube. Fruit ovoid, 15–20 by 10–15 mm, base rounded, at apex tapering into a narrow beak 6–8 mm long, glabrous; red and pulpy when ripe; fruiting pedicel 2–4 mm long. Seeds 1–4, subcircular in outline, thick, 9 –10 by 8 –9 by 4(–5) mm, pale creamy brown, with conspicuously warty-cracked broad margin, faces tuberculate-warty. Distribution — S India; possibly not in Sri Lanka. Habitat & Ecology — In bushes and thickets, over shrubs; altitude 0–800 m. Flowering and fruiting: May to January. Notes — 1. Solena amplexicaulis resembles vegetatively the other 2 species of Solena and Kedrostis foetidissima, and also Adenia wightiana (Passifloraceae). Sterile as well as flowering and fruiting specimens of S. amplexicaulis may be confused also with Kedrostis foetidissima (see introduction and key to the species). 2. Solena amplexicaulis is much more different from the other three taxa of Solena than those are from one another. 2. Solena heterophylla Lour. Solena heterophylla Lour. (1790) 514. — For references and synonyms see under the subspecies.

Plant sparingly pubescent in younger parts, glabrescent, rarely densely hairy; dioecious or monoecious (subsp. napaulensis). Leaves ± ovate, 3–10(–22) cm long, to 15 cm wide, entire, or 3- or 5-angular, or 3- or 5- (long-)lobed to various depth, the lobes broad or (very) narrow, radiating or the lower ones frequently hastate, base deeply but rather narrowly cordate, apex acute(-acuminate), margin minutely or conspicuously sparsely 1–2 mm mucronate-dentate; nerves 3- or 5-palmate, reticulation distinct on both surfaces; glands several or many, small, scattered, most glands near the inser-

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tion of the petiole; petiole 0.5–1(–1.5) cm. Probract 1–5 mm long, caducous. Male inflorescences with a solitary (abortive) flower at the node, co-axillary with 1 (or 2) persistent sub-umbellate or brush-like dense raceme(s), sessile or to 4 cm peduncled, flowers numerous, sparsely hairy, leaving persistent pedicels. Male flowers: pedicel 3–6(–10) mm long, articulate at apex; bract lanceolate or linear, 2–5 mm long, inserted on the pedicel (sometimes towards the apex), somewhat fleshy, with or without glands, persistent or caducous; receptacle-tube 3–4(–5) by 2–3 mm; sepals triangular, 0.2–0.5 mm long, wide apart; petals triangular, 1–2 mm long, finely papillose-hairy (without gland-hairs); filaments 2–3 mm long, glabrous, inserted at or slightly above the base of the receptacle-tube; anthers subglobose, 1–2 mm long, connective broad, convex above, papillose-hairy; thecae oblique or horizontal, curved, forming a nearly closed ring; disc lobes 3 (or 4), papillose-hairy, 1– 2 mm long. Female flowers solitary (rarely a few in a short-shoot), or co-axillary at base of male inflorescence (subsp. napaulensis); pedicel (3–)5–10 mm long; ovary (narrowly-)ellipsoid, c. 10 mm long, short (densely) hairy, with or without scattered or irregular lines of dark-coloured dots; perianth resembling male but slightly larger, ± persistent in young fruit; style c. 2 mm long; stigma, consisting of 3 subglobose long-papillose lobes; staminodes 3 (or 4), elongate, 0.5–1 mm long, glabrous, inserted towards the base of receptacle-tube; disc deeply lobed or consisting of 3 (or 4) free lobes, 1–2 mm long, hairy. Fruit (narrowly-)ellipsoid, 2–5 by 1.5–2.5 cm, velutinous or scabrous, green, paler striped; when ripe yellow or red; pulp yellowish; fruiting pedicel 5–40 mm long. Seeds (5–)10–20, nearly globose or somewhat elongate, little compressed, (5–)6 –7 mm long, c. 5 mm thick, smooth, with a very faint border, grey. Distribution — Two subspecies. Widely distributed, from NE Afghanistan, the Himalayas, through Northern India to S & E China, and SE to Myanmar, Thailand, Indochina, Peninsular Malaysia and Java; not in Sumatra, Borneo. Habitat & Ecology — Prefers a seasonal climate, often in open ± deciduous forest, pine savannah; on granitic as well as on calcareous soils; 0–3300 m altitude. Flowering: March to November; fruiting: April to November. Notes — 1. The forms according to leaf shape, as proposed by Cogniaux (1916), and repeated by Keraudren (1975) have no taxonomic value, although Smitinand on a field label (Thailand) stated that deeply narrow-lobed leaf forms are found in the mountains. 2. In living specimens the leaves are conspicuously pale, glaucous beneath. Unripe fruits are green with irregular whitish stripes, ripe fruits red. KEY TO THE SUBSPECIES

1a. Monoecious (always?). Anther-thecae vertical, sigmoid. Fruit (ovary) with dark dots; fruiting pedicel 20–40 mm long. — Mountainous areas; N India, Nepal, China, Myanmar . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . b. subsp. napaulensis b. Dioecious. Anther-thecae oblique, or (sub)horizontal, semi-circular curved. Fruit (ovary) with paler irregular stripes, rarely with few dots; fruiting pedicel 5–15 mm long. — Widespread, in lowland or montane areas . . . . . . a. subsp. heterophylla



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a. subsp. heterophylla — Fig. 2h, i Solena heterophylla Lour. (1790) 514; (1793) 629; Keraudren (1975) 63, pl. 10, f. 1– 5. — Melothria heterophylla (Lour.) Cogn. (1881) 618; (1916) 121; Backer in Backer & Bakh. (1964) 297 — Type: Lourero s.n. (BM), Cochinchina and China. Bryonia rheedii Blume (1826) 925; Ser. (1828) 306. — Karivia rheedii (Blume) M. Roem. (1846) 45; Miq. (1855) 661. — Type: Blume s.n., barcode: L0127474 (L), Java. Bryonia sagittata Blume (1826) 925; Ser. (1828) 305. — Type: Blume s.n., barcode: L0127475 (L), Java. Zehneria connivens Miq. (1855) 656. — Type: Horsfield s.n. (K), Java. Zehneria hastata Miq. (1855) 656, nom. illegit. — Voucher specimens: Zollinger 669 (L), Horsfield s.n. (K), both Java. Melothria delavayi Cogn. (1916) 124. — Solena delavayi (Cogn.) C.Y. Wu (1984) 341, for the type only, see note 3 under subsp. napaulensis. — Type: Delavay 2009 (lecto, here chosen, P, male and one detached fruit), Yunnan. Zehneria umbellata auct. non (Willd.) Thwaites: C.B. Clarke (1879) 625, p.p.; Ridl. (1922) 851. Solena amplexicaulis auct. non (Lam.) Gandhi: C. Jeffrey (1980b) 14; A.M. Lu & Zhi Y. Zhang (1986) 178, pl. 47, f. 1–9; Grierson (1991) 257, p.p.; S.K. Chen (1995) 322.

Dioecious (always?), with climbing or prostrate shoots. Leaves entire, ovate or hastate, or 3- or 5-angular, or shallowly or deeply 3- (rarely 5-)lobed. Male flowers: anthers with thecae in oblique or (sub)horizontal position, semi-circular, the two thecae forming a nearly closed ring below the broad hairy connective. Female flowers: pedicel 3–5 mm long; ovary hairy, without or with few irregular glabrous bands or blotches. Fruit 2–5 by 1.5–2.5 cm; fruiting pedicel 5–10(–15) mm long. Seeds as described under the species. Distribution — Widespread, as the species. Habitat & Ecology — As the species, but apparently not occurring above 2000 m altitude. Note — A deviating male specimen, Duthie 21434, from Pilhibit District, W Hima­ laya, c. 2000 m altitude, differs from all other specimens by having 1 or 2 long-pedicelled male inflorescences from each node, with many-flowered racemes provided with persistent, relatively large glandular bracts. In general habit it resembles subsp. napaulensis, but the specimen is male (not monoecious) and the thecae (in bud) are curved, not sigmoid. b. subsp. napaulensis (Ser.) W.J. de Wilde & Duyfjes, comb. & stat. nov. — Fig. 1a–c Bryonia napaulensis Ser. (1828) 307. — Zehneria umbellata (Klein ex Willd.) Thwaites var. napaulensis (Ser.) C.B. Clarke (1879) 625 (‘nepalensis’). — Melothria heterophylla (Lour.) Cogn. var. nepalensis (C.B. Clarke) Dhaliwal & M. Sharma (1999) 302. — Type: Wallich s.n. (G, photograph seen), Nepal. ?Bryonia wallichiana Ser. (1828) 309. — Type: Wallich s.n. (G, not seen), Nepal. Solena delavayi auct. non (Cogn.) C.Y. Wu: C.Y. Wu (1984) 341, excl. the type; A.M. Lu & Zhi Y. Zhang (1986) 180, pl. 47, f. 10, 11; S.K. Chen (1995) 324 (see note 3).

Plant monoecious (or dioecious?), shoots often prostrate. Leaves subentire to deeply spreadingly 5-lobed, the middle lobe much longer. Inflorescences either male, with flowers fascicled in a sessile, or a short-peduncled raceme, usually with a co-axillary

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c

a b

c

e

j

h i

k

f

d

g

Fig. 2. a–i: Solena heterophylla Lour. subsp. heterophylla. a, b. Habit of male flowering specimens; c. ditto, a specimen with atypical more or less compound inflorescences resulting from axillary short-shoots; d. ditto, note gland-bearing bracts and bracteoles; e, f. male flower; g. stamens; h. fruit; i. seed. — j, k: Solena umbellata (Klein ex Willd.) W.J. de Wilde & Duyfjes. j. Fruit; k. seed (a: Koorders 41273; b: Zollinger 669; c – g: Garrett 1192; h, i: Whitmore FRI 20266; j: Matthew & Charles RHT 2287; k: Matthew & Charles RHT 51343).



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single rather long-pedicelled female flower, or female flowers solitary at the nodes. Male flowers: as in subsp. heterophylla, but thecae vertical and sigmoid, not forming a nearly closed ring. Female flowers: pedicel 10–25 mm long; ovary hairy, mostly with few or many, glabrous glossy blackish spots. Fruit (broadly) ellipsoid, 3–3.5 by 2–2.5 cm, scabrous-hairy or subglabrous, base and apex ± rounded; fruiting pedicel 20–40 mm long. Seeds c. 7 by 6–6.5 by 3–3.5 mm, grey-brown, smooth, not bordered. Distribution — N India, Nepal, S China (Yunnan), Myanmar. Habitat & Ecology — Mountainous areas at (1000?–)2000–3300 m altitude. Flowering and fruiting: April to August. Notes — 1. The status of S. heterophylla subsp. napaulensis as a subspecies is unclear because in general habit, vegetatively as well as in most flower details, it is very similar to subsp. heterophylla. However, the strikingly S-shaped thecae and the fruit (according to Cogniaux (1916), for Melothria delavayi smaller, and glabrous) distinguish it clearly. More complete material should be studied to ascertain whether the sigmoid thecae is a valid character, or that this character-state may occur occasionally in specimens of local populations within the vast distributional area of S. heterophylla. 2. Specimens from high altitudes in the Himalayas, with 5-lobed leaves, monoecious flowers, the female flowers long-pedicelled, and ovary and fruit having dark dots, are quite distinct. However, several collections, partly from lower areas and often incomplete, look similar but may have characters tending to be intermediate between subsp. napaulensis and subsp. heterophylla; see also note 3. A specimen obviously belonging to subsp. napaulensis but with the thecae only weakly sigmoid is Clarke 28348 (K), from Kashmir. 3. In Chinese literature (Wu, 1984; Lu & Zhang, 1986; Chen, 1995) the present taxon is named Solena delavayi, but we think that this name is synonymous with S. heterophylla subsp. heterophylla. In the original description of Melothria delavayi Cogn. the female flowers are erroneously described as situated solitary in the axil of the male raceme (“solitarii in eadem axilla cum masculis”). The anthers are described as oblong, c. 1 mm long, with ciliate loculi. The S-shape of the thecae is neither indicated in this description nor in the preceding key to the species (Cogniaux, 1916: 80). Moreover, on the lectotype-sheet in P (Delavay 2009, a male plant), Cogniaux made a pencil drawing of an opened male flower, depicting the thecae as only somewhat curved and erect, not S-shaped. In most specimens of subsp. napaulensis that we studied, the anthers with the vertical, sigmoid thecae are c. 2 mm long. Contrary to the statements by Cogniaux (1916: 124) and Jeffrey (1980b: 15), the type material Delavay 2009 consists of different elements, viz. female and male flowering plants: in P male flowering with a loose fruit; in K female flowering and fruiting, and with some loose fruits in an envelope. The male material in P is chosen as lectotype. 3. Solena umbellata (Klein ex Willd.) W.J. de Wilde & Duyfjes, comb. nov. — Fig. 2j, k Bryonia umbellata Klein ex Willd. (1805) 618; Ser. (1828) 305; Wight & Arn. (1834) 345; Dalzell & Gibs (1861) 101. — Momordica umbellata (Klein ex Willd.) Roxb. (1832) 710. — Karivia umbellata (Klein ex Willd.) Arn. (1841) 275; Miq. (1855) 661. — Zehneria umbellata (Klein ex Willd.) Thwaites (1859) 125; C.B. Clarke (1879) 625, p.p. — Type: Klein 765 (lecto B-W, here chosen, see notes), S India.

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Bryonia teedonda Roxb. in Willd. (1805) 618, nom. nud., inval. Karivi-valli Rheede (1688) 51, t. 26. Melothria angulata Chakrav. (1952) 899; (1959) 165, pl. IIB, map 81, f. 74. — Type: Malcolmpeth 81 (CAL), S India. Solena amplexicaulis auct. non (Lam.) Gandhi: Matthew (1983) 651, p.p.; (1999) 541; Philcox (1997) 40, p.p.

Creeper or climber, to 3 m, subglabrous; dioecious; roots tuberous. Leaves ovate or narrowly elliptic, 4–14(–20) by 2–10 cm, entire or deeply hastately 3-lobed, adaxially finely scabrous, abaxially with scattered minute gland-hairs, base cordate, apex acute or obtuse, margin remotely dentate; petiole 4–11 mm long. Probract absent. Male inflorescences subsessile or to 7 mm long peduncled, condensed, few- or many-flowered sub-umbellate racemes up to 1 cm long. Male flowers: pedicel 2–8 mm long, articulate at apex; bracts carnose, ± terete, (narrowly-)ellipsoid, 1–2 mm long, apiculate, without glands, inserted on upper half of the pedicel, (sub)persistent; corolla and anthers very much the same as in S. heterophylla, except for glands on petals: receptacle-tube 3–5 by 2–4 mm; sepals triangular, 0.1–0.2 mm long, wide apart; petals triangular, 1–1.2 mm long, with simple papillose hairs and with numerous minute gland-hairs; filaments 2.5–4 mm long, glabrous, inserted at or slightly above the base of the receptacle-tube; anthers subglobose, c. 1.5 mm diam., connective broad, convex, with gland-hairs; thecae curved, forming an oblique or horizontal nearly closed ring; disc lobes 3, coarsely hairy and with scattered gland-hairs, c. 1 mm long. Female flowers similar to those of S. heterophylla: pedicel 5–10 mm long; ovary glabrous. Fruit solitary at the node, (narrowly-)ellipsoid, 3–7 by 1.5–3 cm, at both ends attenuate (acute, not narrowed into a beak), glabrous, smooth, slightly angled; when dry yellowish, with c. 9 faint ribs, when ripe red; fruiting pedicel 10–12(–15) mm long. Seeds 10–20, subcircular in outline, 6.5–7.5 by 5–6.5 by 3–5 mm, the faces light (grey-)brown, and with a broad paler, finely corky-warted and cracked marginal belt, rendering the seed base somewhat emarginate. Distribution — S India, Sri Lanka. Habitat & Ecology — Thickets, roadsides; altitude 0–1500 m. Flowering and fruiting all year round. Notes — 1. Willdenow (1805) presents a number of references to previous authors, the first being “Bryonia umbellata. Klein in litt.”. Our present species is represented in B-W by two collections: one unnumbered collection by Roxburgh, and one collection by Klein, annotated: “765 Bryonia sp. nov. …… Nandaradah”. The latter sheet, with the smaller leaves, has been chosen as lectotype. In K there are two sheets of the Rottler herbarium (Herb. Rottlerianum), one sheet with 3, the other sheet with 8 separately labelled different collections, all representing the present S. umbellata. Among these collections obviously isotype material is present, but the precise assignment of which fragment is an isolectotype needs further consideration. 2. The seed figured by Chakravarty (1952, pl. 1, K) may represent seeds of S. amplexicaulis, because of the tuberculate faces.



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ACKNOWLEDGEMENTS We thank the keepers of the herbaria BK, BKF, BM, BO, K, KEP, L, LA, P, SING for permitting the study of their specimens and Jan van Os (Leiden) for preparing the drawings. The Reverend K.M. Matthew, S.J., FLS, Director of Rapinat Herbarium, Tiruchirapalhi, S India, provided recent herbarium collections, which proved to be instrumental for the understanding of the species concept.

REFERENCES Arnott, G.A.W. 1840. Remarks on the fruit of the natural order Cucurbitaceae. Madras J. Lit. Sci. 12: 48–54. Arnott, G.A.W. 1841. On the Cucurbitaceae. J. Bot. 3: 271–280. Backer C.A. & R.C. Bakhuizen van den Brink Jr. 1964. Flora of Java 1: 292 – 307. Noordhoff, Groningen. Blume, C. 1826. Bijdragen tot de flora van Nederlandsch-Indië 15: 922 – 940. Ter Lands Drukkerij, Batavia. Chakravarty, H.L. 1952. New finds of Indian Cucurbitaceae. J. Bomb. Nat. Hist. Soc. 50: 894–901. Chakravarty, H.L. 1959. Monograph on Indian Cucurbitaceae. Rec. Bot. Surv. India 17: 1– 234. Chen, S.K. 1995. Solena. In: C.Y. Wu, C. Chen & S.K. Chen (eds.), Flora Yunnanica 6: 322 – 324. Science Press, Beijing. Clarke, C.B. 1879. Cucurbitaceae. In: J.D. Hooker, The flora of British India, 2: 604 – 635. Reeve & Co., London. Cogniaux, C.A. 1881. Cucurbitaceae. In: A. & C. de Candolle, Monogr. Phan. Prodr. 3: 325 – 951. Masson, Paris. Cogniaux, C.A. 1916. Cucurbitaceae-Fevilleae et Melothrieae. In: A. Engler, Pflanzenreich 66, iv.275.1: 1–277. Engelmann, Leipzig. Dalzell, N.A. & A. Gibs. 1861. The Bombay flora or short description of all the indigenous plants: 99–103. Education Society’s Press, Byculla. De Lamarck, J.B.A.P.M. 1785. Encyclopédique Méthodique Botanique 1, 2: 496. Panckoucke, Paris. De Loureiro, J. 1790. Flora cochinchinensis 2: 514. Lisbon. De Loureiro, J. 1793. Flora cochinchinensis 2: 629 (ed. 2, edited by C.L. Willdenow). Berlin. De Wilde, W.J.J.O. & B.E.E. Duyfjes. 2003. Revision of Neoalsomitra (Cucurbitaceae). Blumea 48: 99 –121. Dhaliwal, D.S. & M. Sharma. 1999. Flora of Kullu District, Himachal Pradesh. Dehra Dun, India. Gamble, J.S. 1919. Flora of the presidency of Madras 1, 3: 526– 543. Adlard & Son, Ltd, London. Gandhi, K.N. 1976. Cucurbitaceae. In: C.J. Saldanha & D.H. Nicolson (eds.), Flora of Hassan district Karnataka, India: 174–181. Amerind Publishing Co. Pvt. Ltd, New Delhi. Grierson, A.J.C. 1991. Cucurbitaceae. In: A.J. C. Grierson & D.G. Long, Flora of Bhutan 2, 1: 246–270. Royal Botanic Garden, Edinburgh. Jeffrey, C. 1980a. Further notes on Cucurbitaceae: V. The Cucurbitaceae of the Indian subcontinent. Kew Bull. 34: 789 – 809. Jeffrey, C. 1980b. The Cucurbitaceae of Eastern Asia: 1– 60. Roy. Bot. Gard., Kew. Keraudren-Aymonin, M. 1975. Cucurbitacées. In: A. Aubréville & J.-F. Leroy (eds.), Flore du Cambodge, du Laos et du Viêt-nam 15: 1–114. Mus. Nat. d’Hist. Nat. Paris. Lu, A.M. & Zhi Y. Zhang. 1986. Solena. In: A.M. Lu & S.K. Chen (eds.), Fl. Reip. Pop. Sin. 73, 1: 177–180. Science Press, Beijing. Matthew, K.M. 1982. Illustrations on the flora of the Tamilnadu Carnatic 2: 291– 305. Diocesan Press, Madras. Matthew, K.M. 1983. The flora of the Tamilnadu Carnatic 1: 633– 656. Diocesan Press, Madras. Matthew, K.M. 1999. The flora of the Palni Hills, South India 1: 529 – 545. SCTP Offset Press, Christianpet, Vellore. Miquel, F. A.W. 1855. Flora Indiae Batavae 1, 1: 652– 683. Van der Post, Amsterdam.

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Pax, F. 1889. Cucurbitaceae. In: A. Engler & K. Prantl, Die natürlichen Pflanzenfamilien iv.5: 9 – 39. Engelmann, Leipzig. Philcox, D. 1997. Cucurbitaceae. In: M.D. Dassanayake (ed.), A revised handbook to the flora of Ceylon 11: 8–46. Balkema, Rotterdam. Ridley, H.N. 1922. The flora of the Malay Peninsula 1: 842– 853. Reeve & Co., London. Roemer, M.J. 1846. Peponiferarum. Syn. Monogr. 2: 1–118. Landes-Industrie-Comptoirs, Weimar. Roxburgh, W. 1832. Flora Indica 3: 701–728. Thacker & Co., Calcutta. Seringe, N.C. 1828. Cucurbitaceae. In: A.P. de Candolle, Prodromus systematis regni vegetabilis naturalis 3: 297–320. Treuttel & Würtz, Paris. Thwaites, G.H.K. 1859. Enumeratio plantarum Zeylaniae: 125–128. Dulau & Co., London. Von Rheede tot Draakestein, H.A. 1688. Hortus indicus malabaricus 8: t. 26. Amsterdam. Wight, R. 1841. Icones plantarum Indiae orientalis, or figures of Indian plants: pl. 496–507. Pharoah, Madras. Wight, R. & G.A.W. Arnott. 1834. Prodromus florae Peninsulae Indiae Orientalis 1: 340 – 351. Parbury, Allen & Co., London. Willdenow, C.L. 1805. Caroli a Linné Species Plantarum, tomus 4: 598– 627. Nauk, Berlin. Wu, C.Y. 1984. Index Florae Yunnanensis. Institutum Botanicum Kunmingense Academiae Sinicae Edita 1: 334–348. The People’s Publishing House, Yunnan.

IDENTIFICATION LIST

1 2a 2b 3

= = = =

Solena amplexicaulis (Lam.) Gandhi Solena heterophylla Lour. subsp. heterophylla Solena heterophylla Lour. subsp. napaulensis (Ser.) W.J. de Wilde & Duyfjes Solena umbellata (Klein ex Willd.) W.J. de Wilde & Duyfjes

Anderson 537: 2a — Anglade 1092: 3. Bakhuizen van den Brink 607: 2a — Barber 1662: 3 — Beddome 3295: 3 — Blume s.n., barcode: L0127474 (type): 2a; s.n., barcode L0127475 (type): 2a — Bourne July 1897: 3; 30 Oct. 1899: 3; 141: 3; 367: 3; 463: 3; 452: 3; 2628: 3 — Bowes Lyon 2304: 2a. Chantanamuck 134: 2a — Chatterjee 2638: 2a — Chen His Cheng 1016: 2a — Chung 1686: 2a; 1746: 2a; 2199: 2a; 2221: 2a — Clarke 28348: 2b — J. & M.S. Clemens 3670: 2a — Collett 363: 2b; 629: 2b; 770: 2b — Cramer 4791: 3. Dalhousie 01-07-1831: 2b — D’Alleizette May 1903: 2a; April 1908: 2a; June 1909: 2a — De Wilde & Duyfjes 22162: 2a; 22163: 2a — Delavay 2009 (fruit, lecto P; K): 2a — Diraviados RHT 27866: 1 — Du Pasquier in Pételot 1873: 2a — Duport 81: 2a — Duthie 09-08-1899: 2a; 21434: 2a. Einarson, Skärby & Wetterhall 2589: 2b — Evrard 2145: 2a. Forrest 22085: 2a — Fosberg & Sachet 52990: 3 — Fraser 36: 3 — Fukuoka & Ito T34842: 2a. Gallatly 686: 2a — Gamble 4457A: 2b; 4822A: 2b; 9624: 2a; 9721: 2a; 14819: 3 — Gardner 313: 3; 314: 3; 542: 2b — Garrett 1192: 2a — Geesink, Phanichapol & Santisuk 6021: 2a — Grey-Wilson & Phillips 329: 2b — Griffith 752: 2a. Hepper 4517: 3 — Hohenacker 1506: 3 — How 70799: 2a — Huk s.n., 19-08-1890: 2a — Huq 10369: 2a. Ien 304: 2a — Iwatsuki & Fukuoka T1035: 2a. K’Tung-78 6050: 2a — Keng K2656: 2a — King 23-03-1876: 2a — Koorders 28241: 2a; 41024: 2a; 41273: 2a — Kostermans 510: 2a; 1289: 2a. Larsen 33888: 2a — Larsen, Santisuk & Warncke 3037: 2a — Larsen & Suppee Larsen 34173: 2a — Lawson 188A: 1 — Linsley Gressitt 787: 2a. Madras Herb. 10264: 3; 11579: 3 — Maire 6: 2a; 275: 2a; 3696: 2a — Malcolmpeth 81 (India, seen by Chakravarty, CAL): 3 — Malhotra 50980: 2a — Matthew RHT 23615: 1; RHT 72949: 1 — Matthew & Arockissamy RHT 2562: 1 — Matthew, Britto & Rani RHT 28388: 1; RHT 28777: 1 — Matthew & Charles RHT 2287: 3; RHT 41188: 3; RHT 51343: 3; RHT 51795: 3; RHT 73552: 3 — Matthew & Rajendran RHT 44545: 1; RHT 44789: 1 — Maxwell 71-702: 2a; 75-229: 2a; 75-577: 2a; 87-960: 2a; 88-537: 2a; 88-648: 2a; 91-324: 2a; 93-559: 2a; 93-1044:



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2a; 95-922: 2a; 96-972: 2a — McCosh 209: 2b — Meebold 5870: 2a — Metcalf & Chang 581: 2a — Misra 38085: 2a. Ohashi, Kanai, Ohba & Tateishi 773353: 1; 775434: 2a — Ohba, Tikuchi, Wakabayashi et al. 8580071: 2b. Panigrahi 15142: 2a — Pételot 1085: 2a — Pierre 691: 2a; 4361: 2a — Poilane 17323: 2a; 19774: 2a; 21416: 2a — Polunin 101: 2b; 906: 2b; 987: 2b; 1383: 2b — Polunin, Sykes & Williams 155: 2b; 349: 2b; 889: 3; 2486: 2b; 4191: 2b; 4277: 2b; 4470: 2b; 5437: 2b. Ralaveram July 1845: 1 — Ramaswami 1398 (India, seen by Chakravarty, CAL): 3 — Rao 39147: 2a — Rich 221: 2b — Ritchie 308: 3. Saldanha 14755: 3 — Saulière 18: 3; 70 (India, seen by Chakravarty, CAL): 3; 71 (India, seen by Chakravarty, CAL): 3 — Shimizu, Huto & Chaiglom T9001: 2a — Smitinand & Sleumer 1099: 2a; 1340: 2a — Stainton 4934: 2b — Stainton, Sykes & Williams 381: 2b; 748: 2b; 646: 3; 2926: 3; 5333: 3; 8429: 2a. Tang 10608: 2a — Thompson CP 336: 3 — Thomson 06-06-1849: cf. 2a — Thorel 843: 2a — Thwaites CP 3506: 3 — Tsang Wai Tak 15693: 2a; 16799: 2a; 16803: 2a; 16844: 2a — Tsoung 202: 2a. Van Beusekom, Geesink, Phenklai & Wongwan 3586: 2a; 4508: 2a — Van Beusekom & Santisuk 2908: 2a. Wallich 6705: 2a — Whitmore KEP FRI 20266: 2a — Wight 1121: 1; 1122: 3; 1128 [in K 2 sheets: 1 & 3; in L: 1]; 1129 [in K 4 sheets: 3; in L: 2a]; 1130: 1. Zollinger 669: 2a.